Devonian Period, in geologic time, an interval of the Paleozoic Era that follows the Silurian Period and precedes the Carboniferous Period, spanning between about 419.2 million and 358.9 million years ago. The Devonian Period is sometimes called the “Age of Fishes” because of the diverse, abundant, and, in some cases, bizarre types of these creatures that swam Devonian seas. Forests and the coiled shell-bearing marine organisms known as ammonites first appeared early in the Devonian. Late in the period the first four-legged amphibians appeared, indicating the colonization of land by vertebrates.

During most of the Devonian Period, North America, Greenland, and Europe were united into a single Northern Hemisphere landmass, a minor supercontinent called Laurussia or Euramerica. This union of the paleocontinents of Laurentia (comprising much of North America, Greenland, northwestern Ireland, Scotland, and the Chukotsk Peninsula of northeastern Russia) and Baltica (now most of northern Europe and Scandinavia) occurred near the beginning of the Devonian Period. Extensive terrestrial deposits known as the Old Red Sandstone covered much of its northern area, while widespread marine deposits accumulated on its southern portion. The paleoequator (the site of the Equator at a point in the geological past) passed through North America and through China, which was at that time a separate landmass. South America, Africa, India, Australia, and Antarctica were joined into the Southern Hemisphere continent of Gondwana. Parts of this continent were also often covered by seawater.

An ocean covered approximately 85 percent of the Devonian globe. There is limited evidence of ice caps, and the climate is thought to have been warm and equitable. The oceans experienced episodes of reduced dissolved oxygen levels, which likely caused the extinction of many species, especially marine animals. These extinctions were followed by periods of species diversification, as the descendants of surviving organisms filled in abandoned habitats.

The name of the Devonian Period is derived from the county of Devon, Eng. The English geologist Adam Sedgwick and the Scottish geologist Roderick Murchison proposed the designation in 1839 for the marine rocks they encountered in southwestern England, following the recognition by another British geologist, William Lonsdale, that fossilcorals from Torquay in Devon seemed intermediate in type between those of the Silurian System below and those of the Lower Carboniferous System above. This led to the conclusion that the fossil corals were marine equivalents of the terrestrial Old Red Sandstone rocks already known in Wales and Scotland. The recognition that such major paleogeographic differences existed was a great scientific advance, and it was soon confirmed when Sedgwick and Murchison visited Germany and again when Murchison discovered an intercalation of Devonian marine fossils and Old Red Sandstone fishes near St. Petersburg, Russia. By 1843 American geologist and paleontologist James Hall was able to describe equivalent rocks in eastern North America, but precise correlation with European rocks was not achieved until some years later.

The Devonian environment

Paleogeography

The physical geography of the Devonian can be reconstructed using evidence from paleomagnetism, paleoclimate, paleobiogeography, and tectonic events. Because the paleomagnetic data for the Devonian is conflicting, recent efforts to describe the positions of the continents have concentrated on the rock types associated with particular environments. Such methods focus on the distribution of evaporites, shelf carbonates, and corals because present-day deposits of these types have specific, well-known climatic constraints. Faunal distributions are also employed but to a lesser extent.

The distribution of nonmarine fish and marine invertebrate fossils demonstrates that Europe, Siberia, and the Canadian Arctic islands were linked and formed the bulk of Laurussia. During the Devonian, Asia was composed of many separate microplates that are now joined together. Of these, Siberia and Kazakhstania began fusing during the late Devonian and later joined Laurussia, forming the Ural Mountains along the junction.

There is general agreement that the paleoequator crossed the northern part of Laurussia during the Devonian. Paleomagnetic evidence, however, is not clear, and various positions for the exact placement of the paleoequator have been proposed. Though Laurussia was essentially tropical or subtropical, its climatic zones changed somewhat through the course of the Devonian as this landmass migrated northward during Late Devonian and Early Carboniferous times. Evidence for this movement includes the reduction in evaporitic environments in western Canada and the onset of humid and moist conditions in the area of New York.

The southern continents of today were united into the supercontinent of Gondwana during the Devonian approximately along the lines of their present-day continental shelf boundaries. Establishing the position of Gondwana is more difficult than for Laurussia. Some interpretations favour a wide ocean separating these two large landmasses, but this arrangement is thought to be unlikely because of the remarkable occurrences of similar corals, brachiopods, and ammonites in eastern North America, Morocco, and Spain. Yet, even if these areas were close together, their precise positioning is not certain. Based on the similarity in fossils, some researchers would place North Africa adjacent to the eastern North American seaboard during this period. The late Devonian reef developments in Western Australia suggest a near tropical site for this portion of the southern landmass. The positions of the microcontinents that later came together to form Asia are rather uncertain, but many of them probably were either attached or adjacent to the northern margin of Gondwana and migrated north to fuse with the growing area of Asia at several junctures during the later Phanerozoic Eon.

Paleomagnetic evidence is inconsistent regarding the position of the South Pole. Though some researchers postulate a location in central South America, most favour a position south of central Africa or off its southeast coast. The North Pole was in the ocean.

Paleoclimate

Though most environments present today were represented during the Devonian, evidence of glacial deposits is questionable. It is clear that if polar ice caps did exist, they were very much smaller than they are today. It is thus concluded that Earth was warmer during Devonian time than at present.

Warm and equable climates were common, as shown by the wide distribution of evaporite basins in the Northern Hemisphere, by coal deposits in Arctic Canada and Spitsbergen, and by widespread desert conditions and carbonate reefs. Devonian salt deposits indicative of high evaporation rates, and thus of high temperatures, range from western Canada to Ukraine and Siberia and are found locally in Australia. Evidence of cooler average temperatures is provided by annual tree rings in Archaeopteris trunks from New York state that record seasonal growth patterns characteristic of higher latitudes.

Studies of growth lines on Devonian corals indicate that the Devonian year was longer, about 400 days. The lunar cycle, about 301/2 days, was one day longer than it is now.

Devonian life

A highly varied invertebrate fauna that originated in the preceding Silurian Period continued in the Devonian, and most ecological niches of shallow and deep marine water were exploited. The remarkable proliferation of primitive fishes, which has given the period the name the “Age of Fishes,” occurred in both fresh and marine waters. Derivation of carnivorous fishes from mud-eating forms occurred early in the period, and tetrapods (four-legged land animals) were derived from fishes near the middle of the period. Also remarkable is the rise to dominance of vascular plants. Though groves of trees must have arisen earlier to provide the widespread plant debris noted in Devonian deposits, the first known evidence of in-place forests dates from the Middle Devonian.

Atrypa spinosa, of Middle Devonian age, from Erie county, N.Y.

Courtesy of the Buffalo Museum of Science, Buffalo, N.Y.

Invertebrates

The Devonian invertebrates are essentially of the type established during the Ordovician Period. In nearshore sandy and silty environments, bivalves, burrowing organisms, brachiopods (lamp shells), and simple corals abounded. In offshore environments free from land detritus, biostromes and bioherms flourished, rich in corals, stromatoporoids (large colonial marine organisms similar to hydrozoans), crinoids, brachiopods, trilobites, gastropods, and other forms. In deeper waters, goniatite ammonites (a form of cephalopod), which were one of the few new groups to appear, were abundant. Surface waters were occupied by small dacryoconarids (a shelled marine invertebrate) and by ostracods (mussel shrimp) later in the period. Among the Protozoa, both Foraminifera and Radiolaria were well represented, and sponges were locally abundant.

The corals and stromatoporoids were extremely important for building reef facies. The limestone-reef and forereef facies and biostromal limestones are known in many areas of the world. The corals include tabulate corals, such as Favosites and Alveolites, but especially rugose corals (horn corals), which have been used to establish correlations. Stromatoporoids (a type of sponge with a layered skeleton composed of calcium carbonates) such as Amphipora were common rock builders in the mid-Devonian of the Northern Hemisphere. The twiglike form of Amphipora produces a “spaghetti” or “vermicelli” rock. Elsewhere, only simple corals are frequently found.

Bryozoans (marine moss animals superficially similar to corals) were especially common in shallow shelf seas of the period. Both stony and netted forms occurred, but only the latter, the fenestellids, became important during the period.

The brachiopods (lamp shells) are a group of marine filter-feeding species that bear a resemblance to clams but are not mollusks. Brachiopods were present in a multitude of diverse forms during the Devonian Period. The spire-bearing spiriferoids were perhaps the most common and have been used as index fossils. Two groups of importance emerged: the loop-bearing terebratulids and the spiny mud-dwelling productids. At the same time, a number of groups became extinct, including various orthids and the pentamerids.

Molluscan groups were well represented. The marine clams (bivalves) diversified greatly during the period, especially in the nearshore environments. The earliest freshwater bivalves appeared in the Late Devonian. The gastropods were well diversified, particularly in calcareous (calcium carbonate or limestone) environments, and became even more diversified in later periods. The Scaphopoda (tusk shells) first appeared in the Devonian Period. Another significant Devonian event was the emergence of the ammonites from their still-extant nautiloid ancestors. In the chambered shell of the ammonites, internal septa create elaborate patterns where they join the outer shell. The complexity of these suture patterns culminated in the ammonites of the Mesozoic Era. From their origin (probably in the Emsian Age) the evolution of goniatite ammonites, as well as other ammonites, allows detailed zonal subdivisions to be established until the end of the Cretaceous Period. Devonian goniatites have been found on all continents except Antarctica.

Among the arthropods, the giant eurypterids (sea scorpions) are found in the Old Red Sandstone facies. Some were predatory carnivores and probably lived on fish. The first insect, most likely a collembolan (apterygote), from a group of wingless insects that feed on leaf litter and soil, has been recorded from the Devonian Period of Russia and other areas of Asia. Ostracods (a type of crustacean) were locally very abundant; benthic (bottom-dwelling) forms occur in continental shelf sea deposits, and planktonic (floating) forms occur in the Upper Devonian, where their remains form widespread ostracod-slate facies. Trilobites were well developed in size (some up to 61 cm, or 24 inches, long), variety, and distribution. Nearly all have clearly established Silurian ancestors. The most common were the phacopids, which exhibit a curious trend toward blindness in the Late Devonian. Almost all the diverse Lower Paleozoic trilobite stocks that entered the period were extinct before the close, and only the proetaceans survived into the Carboniferous Period.

Among echinoderms, the holothureans, asteroids, and ophiuroids are known, but they are rare. Crinoids were abundant, including free-living types with grapnel-shaped anchors. The blastoids diversified considerably, but the cystoids did not survive the period.

Vertebrates

Conodonts (recently recognized as toothlike elements of very primitive eel-like vertebrates) are abundant in many Devonian marine facies. Conodonts had perhaps their greatest diversification during the Late Devonian and are of major importance for the correlation of rock layers. More than 40 conodont zones are recognized within the Devonian, and these provide a high-resolution biostraphic framework for the period.

Many groups of Devonian fishes were heavily armoured, and this has led to their good representation in the fossil record. Fish remains are widespread in the Old Red Sandstone rocks of Europe, especially in the Welsh borderland and Scottish areas of Britain; these are mostly associated with freshwater or estuarine deposits. In other areas marine fishes are known, and some of these, such as Dunkleosteus (Dinichthys) from the Upper Devonian Period of Ohio, U.S., may have reached 9 metres (30 feet) in length.

Arctolepis, from the Early Devonian Epoch.

Courtesy of the American Museum of Natural History, New York

The earliest fishes, comprising the agnathans, were without jaws and presumably were mud eaters and scavengers. These types are usually called ostracoderms. Some, such as the osteostracan cephalaspids, had broad, platelike armour of varied form; the brain and nerve structures in some of these are well known. The anaspids also were covered with armour in the form of scales. The heterostracans, which include the oldest known fishes, have an anterior armour basically of upper (dorsal) and lower (ventral) plates; Pteraspis is an example. The Early Devonian saw the entry of jawed forms or gnathostomes, and the armoured forms of these, the placoderms, characterize the epoch. The arthrodires, which had a hinged frontal armour in two portions, and the grotesque antiarchs belong here. The close of the Devonian saw the diminution and extinction of most of these groups, but several other groups continued and have a significant later history.

Sharklike fishes, the chondrichthians, have been found in the Middle Devonian. The bony fishes, or osteichthians of current classification, include the climatioid acanthodeans, which had appeared before the period began, but the lungfishes (Dipnoi), the coelacanths, and the rhipidistians made their first appearance during this time. The last group is thought to have given rise to the four-footed amphibians as well as to all other higher groups of vertebrates.

Devonian lungfish (Dipterus).

Encyclopædia Britannica, Inc.

Plants

It is now known that some supposedly Silurian plants, such as those at Baragwanath, Vic., Australia, are actually from the Early Devonian. The Late Silurian record of Cooksonia fossils of the Czech Republic seems to be the earliest unquestionable evidence of vascular plants. Information on spores provided by palynologists would help determine the antecedents of the Devonian plants.

There was a remarkable initiation of diverse types of vascular plants during the Devonian, and a varied flora was established early in the period. Evidence of algae is common; bryophytes first appear, and charophytes are locally common. Freshwater algae and fungi are known in the Rhynie Chert of Scotland. The first known forests are of late Middle Devonian age.

The Psylotophytopsida is the most primitive group of the pteridophytes (ferns and other seedless vascular plants); this group did not survive the Late Devonian. Cooksonia, Rhynia, and others possessing a naked stem with terminal sporangia (spore cases) belong here. In other members, sporangia were borne laterally but no true leaves were developed, and the branching was often of a primitive dichotomous type. Psylotophytopsids form a basic stock from which other groups apparently evolved. Asteroxylon, which occurs with Rhynia, and other Rhynie plants in the Lower Old Red Sandstone Rhynie Chert of Scotland form a link with the lycopsids by having lateral sporangia and a dense leafy stem. Psylotophytopsids soon gave rise to treelike forms and later to the important lepidodendrids of the Carboniferous flora. Another apparent derivative, the sphenopsids, which has jointed branches, is represented by Hyenia and Pseudobornia. Pteropsids also appeared in the Devonian. Primitive gymnosperms are known, and trunks of Archaeopteris up to 1.8 metres (6 feet) in diameter are present in Upper Devonian deposits of the eastern United States and the Donets Basin of Russia and Ukraine. These trunks apparently were carried by water to their current positions.

The rich record of land plants may be related to the fact that the Old Red Sandstone represents the first widespread record of continental conditions. However, the primitive nature of the stocks seen and the absence of a long earlier record, even of detrital fragments of vascular plants, suggest that the colonization and exploitation of land environments were real Devonian events. Fortuitous finds, such as the silicified flora of the Rhynie Chert and the pyritized tissue from the Upper Devonian of New York, have enabled the intimate anatomy of many of these plants to be elucidated in detail equivalent to that of modern forms.

Faunal realms and migrations

There is a marked similarity in the fauna and flora of the Devonian continental facies the world over. Records from such deposits in China containing Early Devonian genera of the armoured fish Cephalaspis and Pterichthys or the widespread Australian records of Bothriolepis, a Late Devonian antiarch, correspond closely to those of the Old Red Sandstone in Europe. Yet, when studied in more detail, specific differences become apparent. It has been suggested that the Baltic fish succession is so rich that the area must have formed a migration centre. This may be so, but the wide distribution of supposed estuarine and freshwater fishes raises many problems. Many of these can be resolved if the continents were closer together during the Devonian than at present.

Cephalaspis

Courtesy of the trustees of the British Museum (Natural History); photograph, Imitor

The marine life of the Devonian gives little evidence of faunal provinces. It is true that in the Lower Devonian the brachiopodAustralocoelia has been recognized only in the Antarctic, the Falkland Islands, South America, South Africa, and Tasmania and that Australospirifer, Scaphiocoelia, and Pleurothyrella share parts of this distribution. These genera are not known in the marine Lower Devonian of northern continents and seem to establish an “austral” fauna of limited circum-Antarctic distribution at this time (if the southern continents were then united as Gondwana). Elements of this fauna are often called “Malvinokaffric” after the Falkland (Malvinas) Islands and the South African Bokkeveld Beds. At other levels in the Devonian, however, provincial distinctions are not apparent, with the exception of local coral provinces that are distinguishable in areas of Asia.

Extinction events

Throughout the Devonian there were periods of widespread hypoxic or anoxic sedimentation (that is, sedimentary events indicated that little free oxygen or no oxygen at all was dissolved in Devonian seas). Some of these are known to be periods of significant extinction, and all are associated with some faunal anomaly in marine strata. These events are named according to the taxa involved. Some are associated with very wide distribution of certain taxa, such as the Monograptus uniformis, Pinacites jugleri, and Platyclymenia annulata events. The Lower Zlichov Event is associated with the extinction of the graptoloids and the appearance of the coiled cephalopod goniatites. Three events are very significant extinction episodes: the Taghanic Event, which formerly was used to draw the boundary between the Middle and Upper Devonian, was a marked period of extinction for goniatites, corals, and brachiopods; the Kellwasser Event saw the extinction of the beloceratid and manticoceratid goniatite groups, many conodont species, most colonial corals, several groups of trilobites, and the atrypid and pentamerid brachiopods at the Frasnian-Famennian boundary; and the Hangenberg Event saw the extinction of phacopid trilobites, several groups of goniatites, and the unusual late Devonian coiled cephalopods, the clymeniids, at the end of the Famennian Stage.

Earlier, certain writers sought to link these events with thin layers of iridium, characteristic of meteorite or bolide impacts. Evidence of a bolide impact, in the form of possible impact ejecta, has been reported in Middle Devonian deposits and is associated with a pulse of extinction. An impact crater about 65 km (about 40 miles) in diameter, the Siljan structure in Sweden, has been dated to approximately 377 million years ago. This places the impact in the middle of the Frasnian stage (about 372.2 million years ago) and also within the Kellwasser extinction. Nevertheless, the connection between this impact and the Kellwasser Event is still being debated.

A stronger environmental link to Devonian extinctions involves the layers of black shale characteristic of low oxygen conditions. Environmental stress is thought to take place when high global temperatures slow the mixing rate between the ocean’s surface and deeper layers. Bottom waters experience a lowered re-oxygenation rate, which may result in the extinction of many marine species. It is still debated whether these events were caused by climatic extremes caused by an increase in the amount of solar energy from the Sun, an amplified greenhouse effect, or by processes wholly confined to the Earth. For example, greater production of organic matter, perhaps owing to an increased influx of nutrients related to the colonization of landmasses by rooted plants, may have made continental seas more susceptible to anoxia.

There is also evidence that extinctions may be associated with rapid global warming or cooling. Particularly in the Late Devonian, extinction events may relate to periods of abrupt cooling associated with the development of glaciers and the substantial lowering of sea level. It has been argued that patterns of faunal change at the Kellwasser Event are consistent with global cooling.

At present it is not possible to connect Devonian extinctions definitively with any single cause, and, indeed, it is probable that extinctions may record a combination of several stresses.